Boechera falcata (Turcz.) Al-Shehbaz, previously known as Arabis turczaninowii Ledeb., is a herbaceous perennial of the East Siberian, boreal-steppe ecotype. It is the sole species of the diverse genus Boechera found on the Eurasian continent, with all other species endemic to North America and Greenland. Likely migrating from North America to Eastern Siberia via the Bering Land Bridge during the Pleistocene glaciation, B. falcata presents a unique case for genomic study. The genus Boechera is notable for its many allodiploid and triploid apomicts, which have arisen through complex hybridization of sexual species and ecotypes. To date, only the genomes of 2 American Boechera species, B. stricta and B. retrofracta, have been sequenced and analyzed. In this study, we sequenced, assembled to the chromosome level, and analyzed the highly homozygous 189.36 Mb genome of B. falcata (2n = 14). Molecular phylogenetic analysis of nuclear and organelle genomes revealed a high degree of relatedness to North American relatives. Cytogenetic analysis identified all 22 genomic blocks of crucifers, showing that 5 of the 7 B. falcata chromosomes are collinear with their ancestral counterparts, while 2 have undergone inversions. Allelic analysis of the apomixis marker APOLLO gene revealed that B. falcata contains only sex alleles. The availability of the B. falcata genome will advance studies of the evolution and phylogeny of Brassicaceae species and the mechanisms of apomixis, providing a crucial resource for future research in plant genetics and breeding.

• Keywords
• Boechera falcata, Brassicaceae, chloroplast genome, chromosome rearrangements, chromosome-level genome assembly and annotation, comparative chromosome painting, genome structure, molecular phylogeny,
• MeSH
• Brassicaceae * genetics   classification   MeSH
• Chromosomes, Plant genetics   MeSH
• Phylogeny * MeSH
• Genome, Plant * MeSH
• Genomics * methods   MeSH
• Publication type
• Journal Article MeSH

Asexual reproduction through seeds in plants (i.e., apomixis) is a heritable trait, and apomixis- linked loci have been identified in multiple species. However, direct identification of genomic elements is typically hindered as apomixis-linked loci and are commonly found in recombination-suppressed and repetitive regions. Heterochromatinized elements, such as B chromosomes and other supernumerary chromosomal DNA fragments have long been known to be associated with asexuality in both plants and animals and are prime candidate regions for the evolution of multiple apomixis factors controlling the individual elements of apomixis. Here, we examined molecular evolution, gene regulation, and chromosomal location of a male apomeiosis factor (UPG2), a long noncoding RNA gene, in sexual and apomictic Boechera with and without male apomeiosis (i.e., balanced and unbalanced apomicts). We revealed the origin of the gene in the apomixis genome on an apomixis-specific, supernumerary heterochromatic Boechera chromosome (Boe1). The UPG2 is active in the tapetum at male meiosis. We found allele classes specific to apomictic and sexual Boechera accessions and a third class that shares the features of both and points to a convergent transition state. Sex alleles are found only in some of the sexual accessions and have higher nucleotide divergence and lower transcriptional activity compared to apo alleles. These data demonstrate selective pressure to maintain the function of UPG2 for unreduced pollen formation in apomicts as the occasional transmission of the allele from unbalanced apomicts into sexual organisms that lead to pseudogenization and functional decay of copies in sexual organisms.

• Keywords
• Boechera, UPGRADE2, apomixis, gene evolution, heterochromatic chromosome, pollen, supernumerary DNA, tapetum,
• Publication type
• Journal Article MeSH